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OCR for page 105
6 Minerals
A number of inorganic elements are essential for normal
growth and reproduction of animals. Those required in
gram quantities are referred to as macrominerals and this
group includes calcium, phosphorus, sodium, chlorine, po-
tassium, magnesium, and sulfur. The macrominerals are
important structural components of bone and other tissues
and serve as important constituents of body fluids. They
play vital roles in the maintenance of acid-base balance,
osmotic pressure, membrane electric potential and nervous
transmission. Those elements required in milligram or
microgram amounts are referred to as the trace minerals.
This group includes cobalt, copper, iodine, iron, manga-
nese, molybdenum, selenium, zinc, and perhaps chromium
and fluorine. Other elements have been suggested to be
essential based on studies in other species but these are
generally not considered to ever be of practical importance
in dairy cattle. The trace minerals are present in body
tissues in very low concentrations and often serve as com-
ponents of metalloenzymes and enzyme cofactors, or as
components of hormones of the endocrine system. A facto-
rial approach was used to describe the requirements for
both the macro- and trace minerals whenever such an
approach could be supported by research data.
Maintenance requirements as described in this model
will include the endogenous fecal losses and insensible
urinary losses. Though technically not correct, it will also
include losses incurred through sweat. The lactation
requirement will be defined as the concentration of the
mineral in milk multiplied by the 4 percent FCM milk
yield. The pregnancy requirement is defined as the amount
of mineral retained within the reproductive tract (fetus,
uterine contents, and uterus) at each day of gestation. For
most minerals, the requirement of the animal pregnant for
<190 days is small and not considered in the model. The
growth requirement is expressed as the amount of mineral
retained/kg body weight gained and entered into the model
as expected average daily gain (ADG).
The sum of the maintenance, lactation, pregnancy, and
growth requirements is the true requirement of the tissues
for the mineral, and is referred to as the "requirement for
absorbed mineral." The diet must supply this amount to
the tissues. Not all the mineral in a diet is available for
absorption. Where data permitted, the availability of min-
erals from forages, concentrates, and inorganic sources was
assigned an absorption coefficient. The model evaluates
the absorbable mineral content of a diet by determining
the available mineral provided by each constituent of the
diet and comparing the sum of the amount of mineral
available from the diet with the requirement of the animal
for absorbed mineral.
For all minerals considered essential, detrimental effects
on animal performance can be demonstrated from feeding
excessive amounts. Generally, the dietary amount required
for optimal performance is well below amounts found to
be detrimental to performance. However, toxicity from
several of the essential minerals, including fluorine, sele-
nium, molybdenum, and copper are unfortunately prob-
lems that can occur under practical feeding conditions.
The National Research Council (1980) described signs of
toxicosis and the dietary concentrations of minerals that
are considered excessive. Certain elements such as lead,
cadmium, and mercury are discussed because they should
always be considered toxic and are of practical concern
because toxicosis from these elements unfortunately occa-
sionally occurs.
Concentrations of mineral elements in both concentrate
and forage foodstuffs vary greatly (Adams, 1975; Coppock
and Fettman, 1977; Kertz, 19981. Reliable or typical analy-
ses of concentrations of some mineral elements (e.g., chlo-
ride and various micromineral elements) in many foodstuffs
are unavailable (Henry, 1995c). A1SO, concentrations
among samples of the same feed type may be quite variable
depending upon such factors as fertilization and manure
application rates, soil type, and plant species (Butler and
Jones, 19731. Concentrations in byproducts or coproducts
also are variable and influenced by the method of process-
ing to produce the feedstuff. Therefore, laboratory analyses
of feeds for macro- and micromineral element content is
105
OCR for page 106
106 Nutrient Requirements of Dairy CattIe
critically important for precise and accurate diet formula-
tion to meet requirements at least cost. Laboratory analyses
using wet chemistry methods is critical for accurate deter-
mination. Near infrared reflectance spectroscopy (NIRS)
is not reliable (Shenk and Westerhaus, 19941.
Estimates of mean concentrations and of variation (stan-
dard deviations) in mineral element content of many com-
monly used foodstuffs are given in Table 15-1 of this publi-
cation. Compositions of inorganic mineral sources com-
monly used in diet supplementation are presented in
Table 15-3.
MAC RO MINE RALS
Calcium
FUNCTIONS
Extracellular calcium is essential for formation of skeletal
tissues, transmission of nervous tissue impulses, excitation
of skeletal and cardiac muscle contraction, blood clotting,
and as a component of milk. Intracellular calcium, while
1/lO,OOO of the concentration of extracellular calcium, is
involved in the activity of a wide array of enzymes and
serves as an important second messenger conveying infor-
mation from the surface ofthe cell to the interior ofthe cell.
About 98 percent of the calcium in the body is located
within the skeleton where calcium, along with phosphate
anion, serves to provide structural strength and hardness
to bone. The other 2 percent of the calcium in the body
is found primarily in the extracellular fluids of the body.
Normally the concentration of calcium in blood plasma is
2.2 to 2.5 mM (9 to 10 mg/dl, or 4.4 to 5 mEq/L) in the
adult cow, with slightly higher values in calves. Between
40 and 45 percent of total calcium in plasma is bound to
plasma proteins, primarily albumin, and another 5 percent
is bound to organic components of the blood such as citrate
or inorganic elements. From 45 to 50 percent of total
calcium in plasma exists in the ionized, soluble form; the
amount being closer to 50 percent at low blood pH and
closer to 45 percent when blood pH is elevated. The ionized
calcium concentration of the plasma must be maintained
at a relatively constant value of 1 to 1.25 mM to ensure
normal nerve membrane and muscle end plate electric
potential and conductivity, which has forced vertebrates
to evolve an elaborate system to maintain calcium homeo-
stasis. This system attempts to maintain a constant concen-
tration of extracellular calcium concentration by increasing
calcium entry into the extracellular fluids whenever there
is a loss of calcium from the extracellular compartment.
When the loss of calcium exceeds entry, hypocalcemia can
occur and this results in loss of nerve and muscle function,
which can in some instances lead to recumbency and the
clinical condition referred to as milk fever. During vitamin
D intoxication, calcium enters the extracellular compart-
ment faster than it leaves resulting in hypercalcemia, which
can lead to soft tissue deposition of calcium.
CALCIUM HOMEOSTASIS
Calcium leaves the extracellular fluids during bone for-
mation, in digestive secretions, sweat, and urine. An espe-
cially large loss of calcium to milk occurs during lactation
in the cow. Calcium lost via these routes can be replaced
from dietary calcium, from resorption of calcium stored in
bone, or by resorbing a larger portion of the calcium filtered
across the renal glomerulus, i.e., reducing urinary calcium
loss. Whenever the loss of calcium from the extracellular
fluids exceeds the amount of calcium entering the extracel-
lular fluids there is a decrease in the concentration of
calcium in plasma. The parathyroid glands monitor the
concentration of calcium in carotid arterial blood and
secrete parathyroid hormone when they sense a decrease
in blood calcium. Parathyroid hormone immediately
increases renal reabsorption mechanisms for calcium to
reduce the loss of urinary calcium, and will stimulate pro-
cesses to enhance intestinal absorption of calcium and
resorption of calcium from bone.
Ultimately dietary calcium must enter the extracellular
fluids to permit optimal performance of the animal. Cal-
cium absorption can occur by passive transport between
epithelial cells across any portion of the digestive tract
whenever ionized calcium in the digestive fluids directly
over the mucosa exceeds 6 mM (Bronner, 19871. These
concentrations are reached when calves are fed all milk
diets and when cows are given oral calcium drenches for
prevention of hypocalcemia (Goff and Horst, 19931. In
nonruminant species, studies suggest that as much as 50
percent of dietary calcium absorption can be passive (Nel-
lans, 19881. It is unknown how much passive absorption
of calcium occurs from the diets typically fed to dairy cattle
but the diluting effect of the rumen would likely reduce the
degree to which passive calcium absorption would occur.
Active transport of calcium appears to be the major route
for calcium absorption in mature ruminants and this pro-
cess is controlled by 1,25-dihydroxyvitamin D, the hormone
derived from vitamin D. By carefully regulating the amount
of 1,25-dihydroxyvitamin D produced, the amount of
dietary calcium absorbed can be adjusted to maintain a
constant concentration of extracellular calcium (DeLuca,
1979; Bronner, 1987; Wasserman, 19811.
When dietary calcium is insufficient to meet the require-
ments of the animal, calcium will be withdrawn from bone
to maintain a normal concentration of extracellular calcium.
If dietary calcium is severely deficient for a prolonged
period the animal will develop severe osteoporosis to the
point of developing fractures still, because the desire to
maintain a normal concentration of extracellular calcium
OCR for page 107
Minerals 107
is so strong, plasma calcium will only be slightly lower than
normal. A sudden large . 1 r 1 .
1 ~~ ~ ~
increase in loss of calcium from
the extracellular pool can result in acute hypocalcemia
before the calcium homeostatic mechanisms can act. This
is discussed further in the section on milk fever (Chapter 91.
REQUIREMENT FOR ABSORBED CALCIUM
The amount of calcium that must enter the extracellular
compartment for maintenance, growth, pregnancy, and lac-
tation is fairly well known and essentially the same equa-
tions were used to predict these amounts as were used in
the 1989 National Research Council publication Nutrients
Requirements of Dairy Cattle.
Maintenance For maintenance of nonlactating cattle, the
absorbed calcium required is 0.0154 g/kg body weight
(Visek et al., 1953; Hansard et al., 19571. For lactating
animals the maintenance requirement is increased to 0.031
g/kg live BW (Martz et al., 19901. The increase in lactating
cows reflects the impact increased dry matter intake (DMI)
has on intestinal secretion of calcium during digestion.
Growth Growth of cattle requires more calcium when
animals are young and actively accruing bone and less as
they reach mature skeletal size. The Agricultural and Food
Research Council (1991) developed an allometric equation
to describe the calcium requirement of growing calves
which will be adopted in this model. The requirement for
absorbed calcium/kg average daily gain is:
Ca (g/day) = (9.83 x (MW022) x (BW-0221) x WG
where MW = expected mature live body weight (kg), BW
= current body weight, and WG = weight gain.
Pregnancy The developing fetus requires a negligible
amount of calcium until the last trimester of pregnancy
(after day 190 of pregnancy), when the fetal skeleton begins
to become calcified. Fetal skeletal calcification is especially
great in the last weeks before parturition. The absorbed
calcium required to meet the demands of the uterus and
conceptus is best described by the exponential equation
of House and Bell (House and Bell, 1993) for any given
day of gestation beyond day 190 as:
Ca (g/day) = 0.02456 e`00558~-00000
— 0.02456 e(005581-0.00007~-~_~)
where t = day of gestation.
Lactation The amount of calcium/kg milk produced var-
ies slightly with the amount of protein in the milk which
in turn varies with breed. The absorbed calcium required/
kg milk produced is 1.22 g for Holstein cows, 1.45 g for
Jersey cows, and 1.37 g for other breeds. Cows require
about 2.1 g absorbed C a/kg of colostrum produced.
CALCIUM ABSORPTION COEFFICIENT
The amount of calcium that must be fed to meet the
requirement for absorbed calcium is dependent on the
availability of calcium from the foodstuffs and inorganic
calcium sources in the diet, and the efficiency of intestinal
calcium absorption in the animal being fed. The amount
of calcium absorbed from the diet will generally equal the
requirement of the body for calcium if the diet contains
enough available calcium. The proportion of dietary cal-
cium absorbed will decrease as dietary calcium increases
above requirement of the tissues for absorbed calcium. To
truly determine the efficiency of absorption of calcium
from a feedstuff, the animals being tested should be fed
less total dietary calcium than the amount of absorbed
calcium required to meet their needs. This will ensure
that intestinal calcium absorption mechanisms are fully
activated so that the animal will absorb all the calcium
from the foodstuff that it possibly can. Few studies fulfill
this requirement; thus, it is likely that the published data
have underestimated the availability of calcium in many
cases. Previous National Research Council (1978, 1989)
publications have determined a single efficiency of absorp-
tion of dietary calcium regardless of the source of calcium.
This absorption coefficient was 0.38 in the 1989 Nutrient
Requirements of Dairy Cattle and 0.45 in the 1978 Nutrient
Requirements of Dairy Cattle based on the average propor-
tion of calcium absorbed during a variety of trials. The
coefficient was reduced in the 1989 Nutrient Requirements
of Dairy Cattle partly in response to reports that cows in
early lactation were less able to utilize dietary calcium
(Van's Klooster, 1976; Ramberg, 1974) making use of a
lower coefficient for calcium absorption more prudent. The
decision to utilize 0.38 as the calcium absorption coefficient
was based largely on a summary of 11 experiments with
lactating dairy cows in which the average percentage of
dietary calcium absorbed was 38 (Hibbs and Conrad, 19831.
In the majority of these 11 experiments, the cows were
fed diets supplying calcium well in excess of their needs
placing the cows in positive calcium balance by as much
as 20 to 40 g/day. In 3 of the experiments, the cows were
in negative calcium balance and the percentage of dietary
calcium absorbed was still below 40 percent. In those
experiments, alfalfa and/or brome hay were supplying the
dietary calcium. The French Institut National de la Recher-
che Agronomique (1989) used 30 to 35 percent as an esti-
mate of efficiency of absorption for dietary calcium using
similar logic. The 1996 Nutrient Requirements of Beef Cat-
tle utilized 50 percent as the calcium absorption co-
eff~cient. The 1980 United Kingdom Agricultural Research
Council (Agricultural Research Council, 1980) chose 68
OCR for page 108
108 Nutrient Requirements of Dairy CattIe
percent as the coefficient of absorption for calcium; a coef-
ficient considerably higher than the estimate of other
groups that had examined dietary calcium requirements
of cattle. This number is based on a model that predicts
calcium is absorbed by dairy cattle according to need. Using
data from a variety of balance studies in which a substantial
number of lactating cows was included, the observed eff~-
ciency of absorption of calcium (diets utilizing foodstuff
and mineral sources of calcium) reached a plateau of about
68 percent. Concern over the validity of this coefficient
prompted the Agricultural Research Council (1980) to
form a second committee to review the 1980 recommenda-
tion for calcium. This technical committee agreed with the
use of 68 percent as an estimate of the absorption coeff~-
cient to be used in calculating dietary calcium requirements
of cattle (AFRC, 19911.
A single coefficient is inappropriate and in this model
the coefficient for calcium absorption will be based on the
sources of calcium used in the diet. Unfortunately, our
knowledge of the efficiency of absorption of calcium from
individual foodstuffs is limited. Martz et al. (1990) fed
lactating dairy cows two diets with no added mineral
sources of calcium in which alfalfa supplied nearly all of
the dietary calcium. One diet was 33 percent alfalfa, 39
percent hominy grits and 21.5 percent corn cobs; the sec-
ond diet was 24 percent alfalfa, 41.5 percent corn silage
and 29 percent hominy grits. The diets contained more
calcium than suggested by the 1978 National Research
Council Nutrient Requirements of Dairy Cattle publication
and less than suggested by the 1989 National Research
Council Nutrient Requirements of Dairy Cattle. True
absorption of calcium from alfalfa, corrected for endoge-
nous fecal calcium loss, was 25 percent; whereas, from the
alfalfa-corn silage ration 42 percent of calcium was truly
absorbed. Ward et al. (1972) estimated the efficiency of
absorption of calcium from alfalfa ranged from 31 to 41
percent. About 20 to 30 percent of calcium within plants
is bound to oxalate which is relatively unavailable to the
ruminant (Ward et al., 19791. Studies of Hibbs and Conrad
(1983) where cows were in negative calcium balance and
were fed only alfalfa or alfalfa/brome diets fit the criteria
of determining calcium absorption in animals that are being
fed less calcium than they require and in these studies the
efficiency of absorption of calcium from alfalfa ranged from
8 to 37 percent. Because alfalfa is a major contributor of
calcium in dairy rations, absorption of calcium from alfalfa
is used as an estimate of efficiency of absorption of calcium
from forages in general. An efficiency of absorption of 30
percent is used in the model for calcium from forages.
Availability of calcium from grains and concentrates has
not been determined in ruminants. In nonruminant ani-
mals, the availability of calcium from concentrates gener-
ally is less than the availability of calcium from an inorganic
source such as calcium carbonate (Soares, 19951. The pres-
ence of phytate is felt to be a factor impairing absorption
in nonruminants. This is not a factor in ruminants. Because
oxalate is not as likely in concentrate foodstuffs the propor-
tion of calcium available should be greater than 30 percent.
It is possible that it may be comparable to that of the
mineral sources of calcium. However the current model
uses 60 percent as a conservative estimate of the proportion
of calcium available from concentrate foodstuffs based in
part on an assumption that the availability is not as high
as from calcium carbonate. Efficiency of absorption of
calcium from foodstuffs that are not forages (e.g., concen-
trates) was set at 60 percent.
Most non-forage foodstuffs will contain only small
amounts of calcium. However, a notable exception is the
calcium soap of palm oil fatty acids, which can be 7 to 9
percent calcium. The fat of this product is approximately 80
percent digestible, and digestion can only occur following
dissociation of the calcium from the palmitate in the small
intestine. This also implies that 80 percent of the calcium
in this feed ingredient is available for absorption. This is
in contrast to the work of Oltjen (1975) which suggested
that formation of calcium soaps within the rumen impaired
calcium absorption necessitating an increase in diet calcium
when fat was added to a ration. No effect of added fat
on apparent absorption of calcium was observed in the
experiments of Rahnema et al. (19941. The model does
not include a factor to increase dietary calcium when fat
is added to the diet. There may be a need to increase diet
magnesium when fat is added to the diet as magnesium
must be soluble in the rumen to be absorbed. Since hypo-
magnesemia can affect calcium metabolism (see Chapter
9) there is an effect of diet fat on calcium metabolism but
it is not overcome by adding calcium to the ration.
Calcium within mineral supplements is generally more
available than calcium in forages and common foodstuffs
(Hansard et al., 19571. Theoretically the factor limiting
mineral calcium absorption is the solubility of the calcium
from the mineral source. Calcium chloride represents a
source of highly soluble calcium. When 45CaCl was used
as a source of radioactive tracer for calcium absorption
studies it was absorbed with >95 percent efficiency in
young calves (Hansard et al., 19541. Calcium chloride is
assigned an efficiency of absorption coefficient of 95 per-
cent. Estimates of the efficiency of absorption of calcium
from calcium carbonate range from 40 percent or 51 per-
cent (Hansard et al., 1957) or up to 85 percent (Goetsch
and Owens, 19851. Unfortunately, these studies were con-
ducted using steers, with very low requirements for
absorbed calcium. The studies of Hansard et al. (1957)
demonstrate that calcium chloride is between 1.2 and 1.32
times more absorbable than calcium carbonate. Therefore,
the efficiency of absorption of calcium from calcium car-
bonate is designated to be 75 percent. The absorption of
calcium from various mineral sources is often compared
OCR for page 109
Minerals 109
to the efficiency of absorption of calcium from calcium
carbonate. Table 15-4 lists a number of common mineral
sources of calcium (including bone meal) and an estimate
of the efficiency of absorption of calcium in each source,
using data summarized by Soares (1995a) and based on
the efficiency of absorption relative to calcium carbonate.
The calcium from limestone generally is slightly less avail-
able than from pure calcium carbonate and has been
assigned an efficiency of absorption coefficient of 70
percent.
EFFECTS OF PHYSIOLOGIC STATE
The amount of available calcium that will actually be
absorbed varies with the physiologic state of the animal.
Hansard et al. (1954) and Horst et al. (1978) reported that
the efficiency of absorption of calcium decreases as animals
age. Young animals absorb calcium very efficiently and
very old animals absorb calcium poorly. As animals age,
there is a decline in vitamin D receptors in the intestinal
tract (Horst et al., 1990), which is thought to reduce the
ability to respond to 1,25-dihydroxyvitamin D. From the
data of Hansard et al. (1954), the difference in efficiency
of calcium absorption in beef steers from 1 to 6 years of
age is nearly negligible. Age was not included as a factor
to adjust dietary calcium requirement in cattle >200 kg
body weight. The absorption coefficient for calcium from
diets normally fed to calves is high and will be considered
to be 90 percent for all calves <100 kg body weight (see
calf section, Chapter 101.
In early lactation nearly all cows are in negative calcium
balance (Ellenberger et al., 1931; Ender et al.1971; Ramb-
erg, 19741. As feed intake increases and calcium intake
increases most cows go into positive calcium balance about
6 to 8 weeks into lactation (Hibbs and Conrad, 1983;
Ellenberger et al., 19311. Cows in the first 10 days of
lactation are at greatest risk of being in negative calcium
balance (Ramberg, 1974) and some are subclinically hypo-
calcemic throughout this period (Goff et al., 19961. Ram-
berg (1974) reported that the rate of entry of calcium into
the extracellular fluid pool from the intestine increased
about 1.55-fold from the day before parturition until 10
days in milk. Thereafter, the rate of entry of calcium into
the extracellular pool from the intestine was not increased
any further. Van't Klooster (1976) demonstrated that cal-
cium absorption increased from 22 percent in late gestation
to 36 percent by day 8 of lactation after which it remained
relatively constant. This represented a 1.6-fold increase in
efficiency of calcium absorption over this 8-day period.
Regression analysis of data of Ward et al. (1972) predicted
that cows need to be fed 5 g C a/kg milk in early lactation
to avoid negative calcium balance. However, there was no
evidence to demonstrate that negative calcium balance in
early lactation was detrimental to the cow provided the
concentration of calcium in plasma remained normal, i.e.,
lactational osteoporosis ensures adequate entry of calcium
from bone into the extracellular calcium pool. During lacta-
tional osteoporosis, data of Ellenberger et al. (1931) suggest
800 to 1300 g of calcium are removed from bone to support
milk production during early lactation and this calcium is
restored to bone during the last 20 to 30 weeks of lactation
and the dry period. This could increase the requirement
for absorbed calcium in later lactation by as much as 8 g/d
to rebuild bone lost during early lactation. No calcium
requirement for rebuilding bone is included in the model.
The effects of calcium-to-phosphorus ratio on absorption
of calcium and phosphorus was once felt to be important
but recent data suggest that the calcium: phosphorus ratio
is not critical, unless the ratio is >7:1 or <1:1 (Miller,
1983a; Agricultural Research Council, 19801.
CALCIUM DEFICIENCY
A deficiency of dietary calcium in young animals leads
to a failure to mineralize new bone and contributes to
retarded growth. Rickets is more commonly caused by a
deficiency of vitamin D or phosphorus but a deficiency of
calcium can contribute to rickets as well. In older animals a
deficiency of dietary calcium forces the animal to withdraw
calcium from bone for homeostasis of the extracellular
fluids. This causes osteoporosis and osteomalacia in the
bones, which makes the bone prone to spontaneous frac-
tures. The concentration of calcium in milk is not altered
even during a severe dietary deficiency of calcium (Becker
et al., 19331.
EXCESS DIETARY CALCIUM
Feeding excessive dietary calcium is generally not associ-
ated with any specific toxicity. Dietary concentrations of
calcium >1 percent have been associated with reduced
DMI and lower performance (Miller, 1983a) but diets as
high as 1.8 percent calcium have been fed with no apparent
problems for nonlactating dairy cows (Beede et al., 19911.
Feeding excessive calcium could interfere with trace min-
eral absorption (especially zinc) and replaces energy or
protein the animal might better utilize for increased pro-
duction. Feeding calcium in excess of requirements has
been suggested to improve performance, especially when
cows are fed corn silage diets. Because calcium is a strong
cation, addition of calcium carbonate to diets above that
required to meet absorbed calcium needs may be providing
a rumen alkalinizing effect to enhance performance.
Phosphorus
Of all dietary essential mineral elements for dairy ani-
mals, phosphorus represents the greatest potential risk if
OCR for page 110
110 Nutrient Requirements of Dairy Cattle
excess is released into the environment contaminating sur-
face waters and causing eutrophication. Accurate and pre-
cise management of phosphorus nutrition is crucial to opti-
mize performance and health of dairy animals, and to
minimize phosphorus excretion.
PHYSIOLOGIC ROLES
Phosphorus has more known biologic functions than any
other mineral element. About 80 percent of phosphorus
in the body is found in bones and teeth. It is present in
bone, along with calcium, principally as apatite salts, and
as calcium phosphate. It is located in every cell of the body
and almost all energy transactions involve formation or
breaking of high-energybonds that link oxides of phosphate
to carbon or to carbon-nitrogen compounds (such as adeno-
sine triphosphate, ATP). Phosphorus also is intimately
involved in acid-base buffer systems of blood and other
bodily fluids, in cell differentiation, and is a component of
cell walls and cell contents as phospholipids, phosphopro-
teins, and nucleic acids.
Phosphorus concentrations in blood plasma normally are
1.3 to 2.6 mmol/L (4 to 8 mg/dl; 6 to 8 mg/dl for growing
cattle and 4 to 6 mg/dl for adult animals). About 1 to 2 g
circulate as inorganic phosphate in blood plasma of a 600-kg
animal. Because of greater concentrations in erythrocytes,
whole blood contains 6 to 8 times as much phosphorous
as plasma. About 5 to 8 g are present in the extracellular
pool of a 600-kg cow. The intracellular concentration of
phosphorus is about 25 mmol/L (78 mg/dl), and total intra-
cellular phosphorus is about 155 ~ in a 600-k~ cow
(Goff, 1998a).
_, , ,
O O
Phosphorus also is required by ruminal microorganisms
for digestion of cellulose (Burroughs et al., 1951) and syn-
thesis of microbial protein (Breves and Schroder, 19911.
Durand and Komisarczuk (1988) recommended that avail-
able phosphorus (from dietary sources and salivary recy-
cling) within the rumen should be at least 5 g/kg of organic
matter digested to optimize degradation of cell walls from
feeds by microbes. When cattle were fed 0.12 percent
dietary phosphorus, ruminal fluid concentration was over
200 mg phosphorus/L, considerably greater than the 20 to
80 mg of phosphorus/L needed for maximum cellulose
digestion in vitro (Hall et al., 1961; Chicco et al., 19651.
This concentration typically is achieved in cattle by salivary
recycling of phosphorus and from diets adequate to meet
the animal's requirement.
PHOSPHORUS UTILIZATION AND HOMEOSTASIS
Net absorption of phosphorus occurs mainly in the small
intestine (Grace et al., 1974; Reinhardt et al., 19881. Only
small amounts are absorbed from the rumen, omasum, and
abomasum. However, little is known about mechanisms
and regulation of absorption anterior to the small intestine
(Breves and Schroder, 19911. Absorption is thought to
occur mainly in the duodenum and jejunum (Care et al.,
1980; Scott et al., 19841. Unlike absorption of calcium,
absorption of phosphorus is in direct relation to supply of
potentially absorbable phosphorus in the lumen of the
small intestine (Care et al., 19801. Presumably, as in nonru-
minants, absorption occurs via two distinct mechanisms. A
saturable vitamin D-dependent active transport system,
separate and distinct from the active transport mechanism
for Ca, is operative when animals are fed low phosphorus-
containing diets. Synthesis of 1,25-dihydroxyvitamin D can
be stimulated when blood phosphorus is very low resulting
in more efficient absorption (Horst, 19861. Passive absorp-
tion predominates when normal to large amounts of poten-
tially absorbable phosphorus are consumed, and absorption
is related directly to the amount in the lumen of the small
intestine and to concentrations in blood plasma (Wasser-
man and Taylor, 19761.
Absorbed phosphorus may be retained or secreted (e.g.,
in milk) for productive functions or secreted into the lumen
of the digestive tract for reabsorption or excretion in feces.
Homeostasis of phosphorus is maintained predominantly
by salivary recycling and endogenous fecal excretion, which
are related directly to the amount of dietary phosphorus
consumed and absorbed. Concentration of phosphorus in
saliva can be 4 to 5 times of that in blood plasma. In cows,
between 30 and 90 g of phosphorus is secreted daily into
saliva (Reinhardt et al., 1988; Scott, 19881. Almost all phos-
phorus in saliva is inorganic (Reinhardt et al., 1988), and
the amount secreted appears to be regulated by parathyroid
hormone (Wasserman, 19811. Inorganic salivary phospho-
rus is absorbed across the intestine with equal or greater
efficiency than dietary phosphorus (Challa et al., 19891.
REQUIREMENT FOR ABSORBED PHOSPHORUS
For the model, the requirement for absorbed phospho-
rus was factorially derived by summing estimates of true
requirements for maintenance, growth, pregnancy, and
lactation.
Maintenance Typically, 95 to 98 percent of total phospho-
rus excretion is in feces. Three fractions are present that
of dietary origin unavailable for absorption or not absorbed,
that of endogenous origin which is inevitably excreted
(inevitable fecal loss), and that of endogenous origin which
is excreted to maintain homeostasis (representing phospho-
rus absorbed by the intestine in excess of the need to
maintain normal blood phosphorus). By definition, the
maintenance requirement of phosphorus is the endoge-
nous fecal loss (inevitable fecal loss) when phosphorus
supply is just below or just meets the true requirement.
In the past, the maintenance requirement was expressed
OCR for page 111
Minerals 1 1 1
as a function of body weight (National Research Council,
1989a), based on fecal phosphorus excretion data extrapo-
lated to zero phosphorus intake (Agricultural Research
Council, 19801. This was later determined to be an inappro-
priate approach (Agricultural and Food Research Council,
19911. Other workers suggested that inevitable fecal loss
in ruminants was a function of total fecal dry matter (DM)
excretion (Conrad et al., 1956; Preston and Pfander, 1964),
which reflects the role of the salivary glands in phosphorus
metabolism. It follows, therefore, that inevitable fecal loss
of phosphorus also is related to DMI. The Agricultural and
Food Research Council (1991) hypothesized that inevitable
fecal loss of phosphorus is determined mainly by DMI,
and not by live body weight. New research was available
with cattle illustrating that a conceptually more sound and
repeatable approach than expression as a function of body
weight is to express maintenance requirement as a function
of DMI when, by definition, dietary phosphorus is fed and
absorbed very near the true requirement.
Part of the maintenance requirement for absorbed phos-
phorus of the animal is the inevitable fecal loss associated
with microbial cells of the digestive tract which contain
phosphorus and are excreted in feces. It is estimated that
about half of the inevitable fecal loss of phosphorus is
associated with microbial debris, and purines and pyrimi-
dines of nucleic acids. This fraction can vary depending
upon fermentability (fermented organic matter) ofthe diet.
However, sufficient data are lacking to quantify this rela-
tionship accurately (Kirchgessner, 19931.
Klosch et al. (1997) fed growing bulls (228 or 435 kg
BW) diets low (50 percent) or high (80 percent) in concen-
trates and total phosphorus balance was determined. Net
phosphorus retention was <1 g/animal per day, and fecal
phosphorus excretion was not influenced by digestibility
of organic matter consumed, or body weight. Total fecal
phosphorus excretion (phosphorus of dietary origin not
absorbed plus that of endogenous origin that was inevitably
excreted) averaged 1.0 g/kg of DMI. The absorption coeff~-
cient of total dietary phosphorus was assumed to be about
80 percent in the study of Klosch et al. (19971. Therefore,
the absorbed phosphorus requirement for maintenance of
growing animals was set at 0.8 g/kg of DMI in the current
model. An additional 0.002 g/kg BW (Agricultural Research
Council, 1980) of endogenous phosphorus loss from urine
was considered part of the maintenance requirement for
absorbed phosphorus in the model.
Spiekers et al. (1993) fed a low phosphorus (0.21 per-
cent) diet to two groups of lactating dairy cows of similar
BW, but differing in daily milk yield (stage of lactation
effect) and feed intake. For the two groups total phospho-
rus intakes were 37 and 21.5 g/day, respectively; and, phos-
phorus balance was similar and slightly negative, indicating
that animals were fed below or very near the true require-
ment. Total excretion of fecal phosphorus differed between
groups (20.3 versus 13.3 g/cow per day) and was 51 percent
greater per kg of body weight for cows at high versus low
dietary phosphorus. However, calculated as a function of
DMI, excretion of fecal phosphorus was 1.20 and 1.22
g/kg DMI per day for the high and low intake groups,
respectively. It is estimated that the absorption coefficient
of total dietary phosphorus for cows fed very close to the
true requirement is 80 percent. Therefore, in the current
model the maintenance requirement for nonlactating preg-
nant and lactating cows was set at 1.0 g/kg of dietary dry
matter consumed. A small amount of endogenous phospho-
rus is inevitably excreted in urine. To account for this, an
additional 0.002 g/kg BW (Agricultural Research Council,
1980) is considered as part of the maintenance requirement
for absorbed phosphorus in the model.
Growth The requirement for growth is the sum of the
amount of absorbed phosphorus accreted in soft tissues
plus that deposited in skeletal tissue. An accretion of 1.2 g
of phosphorus/kg soft tissue gain was estimated by Agricul-
tural Research Council (1980) and data of Grace (1983)
from lambs confirmed this value. However, the majority
of phosphorus deposition in growing animals is associated
with new bone (hydroxyapatite) growth. Bone contains
120 g of calcium/kg and the theoretic accretion ratio of
calcium-to-phosphorus is about 2.1 g calcium-to-1.0 g
phosphorus (1.6 mol per 1.0 mol). Using this relationship
and the accretion rate in soft tissues, the Agricultural and
Food Research Council (1991) developed an allometric
equation from data in the literature with growing cattle
to describe the requirement for absorbed phosphorus for
growth (g/kg average daily gain):
P (g/day) = (1 2 + (4.635 x MW022) (BW-02211) x WG
where MW = expected mature live body weight (kg),
BW = current body weight, and WG = weight gain.
Because bone is an early maturing component of the
body, the allometric equation reflects declining require-
ment for absorbed phosphorus for growing animals. This
equation was used to define the absorbed phosphorus
requirement for growing dairy cattle. For example in the
model, for an animal with M = 681 kg, the absorbed
phosphorus requirements (g/kg average daily gain) ranges
from 8.3 g at 100 kg live BW (C) to 6.2 g at 500 kg.
Pregnancy Quantitatively the requirement for phospho-
rus for pregnancy is low until the last trimester. New infor-
mation on accretion of phosphorus in conceptuses (fetus,
fetal fluids and membranes, placentomes and uterine tis-
sues) of 18 multiparous Holstein cows slaughtered at vary-
ing times from 190 to 270 days of gestation was available
(House and Bell, 19931. Changes in fetal mass and phos-
phorus content across the sampling period were similar
OCR for page 112
112 Nutrient Requirements of Dairy Cattle
to earlier data (Ellenberger et al., 19501. Therefore, the
requirement for absorbed phosphorus to meet demands
of the conceptus for any day beyond 190 days of gestation
is described in the model by the exponential equation:
absorbed phosphorus (g/d)
= 0. 02743e (o 05527 - o~oooo75 t)t
— 0.02743e (0 05527-0 000075 (t- l))(t-
7
where t = day of gestation (House and Bell, 19931.
Estimates of rates of phosphorus accretion in concep-
tuses of Holstein cows increase from 1.9 g/d at 190 to 5.4
g/d at 280 days of gestation, respectively. This equation
should not be used to predict phosphorus accretion of the
conceptus prior to 190 days of gestation. The phosphorus
requirement of the conceptus at <190 days of gestation
is very small and was set to zero in the model.
Lactation The requirement for absorbed phosphorus
(g per day) for lactation is equal to daily milk yield multi-
plied by the percentage of phosphorus in milk. The phos-
phorus content of milk ranged from 0.083 to 0.085 percent
(Wu et al., 2000), 0.087 to 0.089 percent (Spiekers et al.,
1993), and 0.090 to 0.100 percent (Flynn and Power,19851.
The value of 0.090 percent (0.90 g of phosphorus per kg
of milk) was used to compute requirements for absorbed
phosphorus in the model. This is the same as that used by
the working groups in Scotland and the United Kingdom
(Agricultural and Food Research Council, 1991), France
(Gueguen et al., 1989), and Germany (Kirchgener, 19931.
In the last edition of this publication (National Research
Council, 1989), the requirement of phosphorus for lacta-
tion was adjusted depending upon fat content of milk.
However, the phosphorus in cows' milk is distributed as:
20 percent esterif~ed to casein; 40 percent as colloidal
inorganic calcium phosphate; 30 percent as phosphate ions
in solution; and, only about 10 percent associated with the
lipid fraction (;[enness and Patton, 1959; Renner, 19831.
Therefore, an adjustment based on milk fat content is not
of major quantitative and practical significance in defining
the phosphorus requirement for lactation of dairy cows.
DIETARY REQUIREMENT AND EFFICIENCY
OF ABSORPTION
The dietary requirement is the sum of the requirements
for absorbed phosphorus for maintenance, growth, preg-
nancy, and lactation divided by the absorption coeff~cientts)
for phosphorus from the diet. The absorption coefficient
in the denominator of the factorial equation potentially has
more influence on the final computed dietary requirement
than any of the single or combined requirement values for
absorbed phosphorus. The smaller the absorption coeffi-
cient, the greater will be the calculated dietary require-
ment. In the last edition, an overall absorption coefficient of
50 percent was used (National Research Council, 1989b).
Other working groups established overall values of 58 per-
cent (Agricultural and Food Research Council, 1991), 60
percent (NRLO, 1982), 60 percent (Gueguen et al., 1989),
and 70 percent (Kirchgessner, 19931. As with calcium, a
single overall absorption coefficient was not considered
appropriate for all types of feedstuffs, supplemental min-
eral sources, or diets fed to various classes of dairy animals
because of the known variation in absorption coefficients.
The model evaluates the absorbable phosphorus content
of the diet by determining the phosphorus available for
absorption from each ingredient of the diet and comparing
the sum of total phosphorus in the diet with the require-
ment for absorbed phosphorus of the animal.
To accurately determine the true absorption coeti~c~ent
from a particular foodstuff or mineral source, phosphorus
must be fed in an amount less than the animal's true
requirement. This is to insure maximum efficiency of
absorption of all potentially absorbable phosphorus from
that particular source. A1SO7 especially with phosphorus,
the amount of endogenous phosphorus recycled via saliva
must be taken into account. This is most appropriately
done experimentally by quantifying recycling with a tracer
(e.g., p32) Most studies do not satisfy these experimental
specifications. Thus, the true absorption coefficient is gen-
erally unknown and the value given is an underestimation
of true absorption. Apparent absorption of phosphorus (or
apparent digestibility) determined in many studies is lower
(largely because of copious endogenous fecal excretion)
and not equivalent to the true absorption coefficient. If
apparent absorption estimates are used to compute a
dietary requirement, gross over-estimation results.
Based on available data, absorption coefficients of phos-
phorus used in the model for most foodstuffs commonly
fed to cattle of various physiologic states were: 90 percent
for calves consuming milk or milk replacer; 78 percent for
young ruminating calves 100 to 200 kg body weight. True
absorption coefficients for phosphorus from alfalfa hay or
corn silage were 67 percent or 80 percent, respectively,
for lactating cows yielding about 33.6 kg of 3.5 percent
fat-corrected milk and consuming 21.7 kg DM daily (Martz
et al., 19901. Using a tracer technique, Lotgreen and
Kleiber (1953, 1954) reported the true absorption coeff~-
cient of phosphorus in alfalfa hay fed to lambs ranged from
0.81 to 0.96. In the model, absorption coefficients of 64
percent and 70 percent were used for forages and concen-
trates, respectively.
More complete data are available to estimate absorption
coefficients of various potential supplemental mineral
sources (Table 15-41. These values were tabulated from
Soares (1995b) and Peeler (1972), and other sources in
the literature and used in the model. Those values deter-
mined with ruminants, and especially with cattle, were
given preference whenever possible in tabulation.
OCR for page 113
Minerals 1 13
Dicalcium phosphate (calcium phosphate dibasic) with
a true absorption coefficient of 75 percent in cattle (Tillman
and Brethour, 1958; Challa and Braithwaite, 1988), phos-
phoric acid with true absorption coefficient of 90 percent
in cattle (Tillman and Brethour, 1958), and monosodium
phosphate with a true absorption coefficient of 90 percent
in sheep (Tillman and Brethour, 1958) were taken as refer-
ence standards. The absorption coefficients of phosphorus
in other mineral sources were set based on these reference
standards and data where relative differences in phospho-
rus absorption among these and other sources were esti-
mated in various experiments (Soares, 1995b).
Because sufficient studies with appropriate tracers are
not available to estimate true absorption coefficients for
most foodstuffs fed to lactating dairy cattle, an alternate
approach would be useful. One such approach involves
utilizing experimentally derived phosphorus balance data
and the assumption that an accurate estimate ofthe mainte-
nance requirement for absorbed phosphorus is 1.0 g/kg of
DMI (Spiekers et al., 1993) plus endogenous urine output
(0.002 g/kg BW; Agricultural Research Council, 19801. A
calculated absorption coefficient can be derived as: Etrue
requirement for maintenance (g per day) plus milk phos-
phorus output (g per day) plus phosphorus balance (g per
day)] divided by total phosphorus intake (g per day). The
fecal output value from the actual balance determination
is ignored because it represents unabsorbed dietary phos-
phorus plus excess endogenous phosphorus which has been
recycled to the digestive tract via saliva and excreted in
feces. Using this approach, the calculated absorption coeffi-
cients of phosphorus in mixed diets fed to lactating cows
ranged from 67 to 100 percent (Morse et al., 1992b; Spiek-
ers et al., 1993; Brintrup et al., 1993; Wu et al., 20001. In
each study, two or three different concentrations of dietary
phosphorus were fed. Within each study the calculated
absorption coefficient declined as the dietary phosphorus
concentration increased, as would be expected (Challa et
al., 19891. A1SO, among three studies in which dietary phos-
phorus concentrations (0.39 to 0.42 percent) most closely
supplied the requirement of lactating cows, the calculated
absorption coefficients L67 percent, Brintrup et al. (19931;
74 percent, Morse et al. (1992b); 72 percent, Wu et al.
(20001] were similar to the overall absorption coefficient
(70 percent) set by the German working group (Kirchges-
sner, 19931. In the case of Spiekers et al. (1993), in which
lactating cows were fed diets with 0.21 percent phosphorus
(phosphorus-def~cient diet which resulted in slightly nega-
tive phosphorus balance) the calculated absorption coeff~-
cient was about 100 percent, as would be expected. This
relationship is corroborated by regression of the calculated
absorption coefficients on dietary phosphorus concentra-
tions ranging from 15 to 62 percent, dry basis. Regression
analysis (adjusted for number of experimental observations
per treatment mean) was performed with a data set of 71
treatment means from 20 phosphorus balance trials (Hibbs
and Conrad, 1983; Martz et al., 1990; Morse et al., 1992b;
Spiekers et al., 1993; Brintrup et al., 1993; Wu et al., 2000;
Rodriguez, 19981. The regression equation is: calculated
absorption coefficient= 1.86696 - 5.01238(dietaryphos-
phorus percent) + 5.12286 (dietary phosphorus percent)2;
(r2 = 0 70) Based on the regression equation, the calcu-
lated absorption coefficient was 1.0 with 0.22 percent phos-
phorus and declined to a minimum absorption coefficient
of 0.64 with 0.49 percent dietary phosphorus. All of these
calculated absorption coefficients are greater than that
(0.5) used by the National Research Council (19891.
Efficiency of absorption of phosphorus depends upon a
number of factors: age (or body weight) of the animal;
physiologic state (e.g., nonlactating versus lactating);
amount of DM or phosphorus intake; calcium-to-phospho-
rus ratio; dietary concentrations of aluminum, calcium,
iron, magnesium, manganese, potassium, and fat; intestinal
pH; and, source of phosphorus (e.g., forages, concentrates,
inorganic mineral supplements, and salivary phosphorus)
(Irving, 1964; Peeler, 1972; Agricultural and Food
Research Council, 1991; Soares, 1995b).
EFFECT OF INTAKE OF PHOSPHORUS
Efficiency of absorption of phosphorus declines as intake
of phosphorus increases in cattle (Challa et al., 1989) and
in sheep (Field et al., 1977~. However, over a considerable
range of phosphorus intakes within recommended amounts
the efficiency of absorption (absorption coefficient) from
inorganic sources remained high and relatively constant in
cattle (83 percent; Challa et al., 1989) and in sheep (74
percent; Braithwaite, 1986~. Because salivary phosphorus
typically supplies appreciably more (e.g., at least two-fold
greater amounts) phosphorus to the lumen of the small
intestine than does dietary phosphorus, the efficiency of
absorption of salivary phosphorus is important. Salivary
phosphorus is in the form of inorganic phosphate salts
with sodium and potassium. Over a considerable range
of phosphorus intakes in tracer studies, the absorption
coefficient of salivary endogenous phosphorus recycled to
the small intestine was 68 percent to 81 percent in bull
calves (Challa et al., 1989~. Excessive dietary phosphorus
relative to the requirement reduced the efficiency of
absorption of inorganic or salivary phosphorus (Braith-
waite, 1983,1986; Challa et al., 1989~.
EFFECT OF DIETARY CALCIUM
Effect of increasing dietary calcium on phosphorus
absorption was investigated where dietary calcium-to-phos-
phorus ratios ranged from 0.6 to 3.6 (Field et al., 1983~.
Efficiency of absorption of phosphorus in sheep was
reduced by 18 percent with increasing amounts of calcium;
OCR for page 114
114 Nutrient Requirements of Dairy Cattle
amounts of calcium and phosphorus were within those
amounts recommended by Agricultural Research Council
(19801. At higher than recommended supplemental cal-
cium, greater depression of phosphorus absorption would
be expected (Agricultural and Food Research Council,
19911. Phosphorus deficiency was exacerbated in lambs
fed diets supplying 1.5 times daily requirements for calcium
(Seville and Ternouth, 1981), likely a result of reduced
soluble phosphorus in the digestive tract (Wan-Zahari et
al., 19901.
PHYTATE PHOSPHORUS
About two-thirds or more of phosphorus in cereal grains,
oilseed meals, and grain by-products is bound organically
in phytate; stems and leaves of plants contain very little
phytate phosphorus (Nelson et al., 19761. Phytate phospho-
rus is only slightly available or totally unavailable to non-
ruminants (Soares, 1995b; National Research Council,
19981. However, inherent phytase activity of ruminal
microorganisms renders nearly all of the phytate phospho-
rus available for absorption (Reid et al., 1947; Nelson et
al., 1976; Clark et al., 1986; Morse et al., 1992a; Ingalls
and Okemo, 1994; Herbein et al., 19961.
VARIATION IN PHOSPHORUS CONTENT OF FEEDS
Phosphorus is the most expensive macromineral element
supplemented in diets of dairy cattle. Therefore, laboratory
analyses of feeds for phosphorus content is critically impor-
tant for precise and accurate diet formulation to meet
requirements at least cost. There is considerable variation
in actual phosphorus content within types of forages and
concentrates fed to dairy animals (Adams, 1975; Kertz,
19981. Estimates of variation (standard deviations) in phos-
phorus content of many commonly used foodstuffs are
given in Table 15-1 of this publication.
GROWTH AND MILK YIELD RESPONSES TO VARYING
DIETARY PHOSPHORUS CONCENTRATIONS
In addition to the factorial approach for deriving the
absorbed and dietary requirements, results of feeding trials
in which varying dietary concentrations of phosphorus were
fed to growing calves and lactating cows were evaluated.
GROWING CALVES
Huffman et al. (1933) concluded that 0.20 percent
dietary phosphorus was not sufficient for growth of dairy
heifers from 3 to 18 months of age. Maximum weight gains
of dairy calves from 90 to 125 kg BW occurred when dietary
phosphorus content was 0.24 percent, dry basis (Wise et
al., 19581. However, bone ash content was greater when
1. . . ..
dietary phosphorus was 0.33 percent compared with 0.24
percent, but greater phosphorus intake did not improve
any other performance variables. Noller et al. (1977) found
no differences in BW gain, efficiency of converting feed to
gain, or concentrations of phosphorus in blood of Holstein
heifers gaining between 0.68 to 0.82 kg/head per day when
fed diets containing either 0.22 or 0.32 percent phosphorus.
In a second trial, 0.32 percent compared with 0.22 percent
dietary phosphorus increased concentrations of phospho-
rus in serum, but no differences in weight gain or efficiency
of feed conversion were observed. Increasing dietary phos-
phorus from 0.24 to 0.31 percent (dry basis) increased
DMI, average daily gain, breaking strength of ribs and
tibia, and concentrations of inorganic phosphorus in blood
plasma of dairy calves (Teh et al., 19821. Langer et al.
(1985) compared 0.24, 0.3O, and 0.36 percent dietary phos-
phorus fed to growing calves and found over the 10-week
study that 0.30 percent resulted in maximum feed intake,
average daily gain, and concentrations of phosphorus in
blood plasma; no additional benefits were detected with
0.36 percent phosphorus. Miller et al. (1987) fed diets
containing 0.08, 0.14, 0.2O, or 0.32 percent phosphorus and
concluded, from concentrations of phosphorus in blood
plasma and average daily gains, that at least 0.32 percent
phosphorus was needed for heifers to gain 0.75 kg per day.
Two sources (monoammonium phosphate and dicalcium
phosphate) of phosphorus each used to give three dietary
phosphorus concentrations (0.26, 0.34, and 0.41 percent,
dry basis) were compared with growing dairy calves (;Jack-
son et al., 19881. Increasing dietary phosphorus from 0.26
to 0.34 percent increased feed intake, body weight gain,
concentrations of inorganic phosphorus in blood plasma,
and bending moment of the tibia and rib. Body weight
gain (0.94 kg/head per day) of calves fed 0.34 percent
phosphorus was about 13 percent greater than that of calves
fed 0.26 percent dietary phosphorus. Only plasma concen-
tration of phosphorus was increased further with 0.41 per-
cent phosphorus compared with lower concentrations. All
responses were similar between sources of supplemental
phosphorus. Based on all of these studies, 0.30 to 0.34
percent dietary phosphorus was sufficient for normal blood
concentrations of phosphorus in blood, maximum average
daily gains, and greater bone strength of growing dairy
calves.
LACTATING CATTLE
Research literature was reviewed to find all possible
results characterizing lactational responses to varying
dietary concentrations of phosphorus. Phosphorus is often
fed in greater dietary concentrations than needed to meet
the requirement established in the current model. Is feed-
ing phosphorus in excess of requirement beneficial?
OCR for page 115
Minerals 1 15
Nine studies, with dietary phosphorus concentrations
ranging from 0.24 to 0.65 percent of dietary dry matter,
fed for periods ranging from the first 8 weeks of lactation
to as long as three consecutive lactations, with average milk
yields ranging from 15 to 40/kg per cow per day were
examined to try to answer this question.
Overall, supplying more dietary phosphorus than that
calculated to meet the dietary requirement did not increase
DMI or milk yield in any of the studies. The study of
Kincaid et al. (1981) suggested that increasing dietaryphos-
phorus increased DMI and 3.5 percent fat-corrected milk
yield. However, based on the description of the analysis
of variance in that paper the data were improperly ana-
lyzed, thus invalidating interpretation. In one other study,
feed intake and milk yield were lower for cows fed 0.24
versus 0.32 or 0.42 percent phosphorus (Call et al., 19871.
Within none of the other studies was DMI or milk yield
increased by increasing dietary phosphorus from its lowest
concentration to a higher concentration (Stevens et al.,
1971; Carstairs et al., 1981; Brodison et al., 1989; Brintrup
et al., 1993; Dhiman et al., 1996; Wu and Satter, 2000;
Wu et al., 20001.
Milk fat and protein percentages were not attectect by
concentration of dietary phosphorus in most studies. Milk
protein percentage increased as phosphorus increased
from 0.32 or 0.42 percent compared with 0.24 percent
(Call et al., 19871. Protein content of milk was higher with
0.45 versus 0.35 percent phosphorus in the study of Wu
and Satter (20001. Milk fat percentage was higher in year
1 of the study of Brodison et al. (1989) with 0.44 versus
0.35 percent phosphorus, but lower in the study of Brintrup
et al. (1993) with 0.33 versus 0.39 percent phosphorus.
There were no consistent effects of dietary phosphorus
concentration on milk composition among studies.
Concentrations of phosphorus in blood were evaluated
in seven of the nine studies. The normal concentrations
of inorganic phosphorus in plasma is 4.0 to 6.0 mg/dl for
adult cattle (Goff, 1998a). In only one case among all of
the studies was phosphorus in blood below the normal
range (3.6 mg/dl for cows fed 0.24 percent dietary phospho-
rus; Call et al., 19871; 0.24 percent did not provide the
dietary requirement. In other studies, increasing dietary
phosphorus increased the concentration of phosphorus in
blood within or above the normal range.
The DMI and milk yield of cows during early lactation
were maximized with 0.40 to 0.42 percent dietary phospho-
rus, and greater concentrations (0.50 to 0.52 percent) did
not increase DMI or milk yield (Carstairs et al., 1981; Wu
et al., 20001. Milk yield was not affected by the concentra-
tion of the phosphorus in the diet during the first month,
but from week 5 through 12 of lactation, it tended to
be greater with 0.40 percent compared with 0.50 percent
phosphorus (Carstairs et al., 19811. For the entire 84-d
treatment period, cows fed 0.40 percent phosphorus pro-
duced 8 percent more milk than those fed 0.50 percent
phosphorus. Feeding 0.42 percent phosphorus to high
yielding cows during the first 8 weeks of lactation maxim-
ized milk production, and resulted in positive phosphorus
balance and normal concentrations of phosphorus concen-
trations in blood serum (Wu et al., 20001.
Based on results of nine studies, a concentration in the
range of 0.32 to 0.42 percent phosphorus for the entire
lactation was sufficient, depending upon milk production
potential of the cows and nutrition supplied. No benefits
on lactational performance of dietary concentrations >0.42
percent phosphorus were reported in any short- or long-
term studies which were properly analyzed.
Daily dietary requirement determined by the factorial
method is expressed as g per cow per day, and not as a
percentage of the diet. Therefore, supplying the require-
ment requires a reasonably accurate estimate of actual
DMI.
FREE-CHOICE PHOSPHORUS
Coppock et al. (1972, 1975) studied the practice of free-
choice feeding of phosphorus-containing supplements to
dairy heifers and lactating cows to meet requirements when
diets were low or marginally deficient in phosphorus or
calcium. With heifers there was little relationship between
need for the mineral elements and free-choice consump-
tion of dicalcium phosphate or defluorinated phosphate.
For lactating cows offered basal diets providing phosphorus
and calcium below requirements for 9 and 12 weeks, there
was no evidence that cows consumed dicalcium phosphate
to correct the deficiency or that appetite for phosphorus
and calcium supplements coincided with the animals' nutri-
tional requirements.
PHOSPHORUS DEFICIENCY
Detailed description of occurrence, etiology, clinical
pathology, diagnosis, treatment, and prevention of phos-
phorus deficiency in ruminants has been described by Goff
(1998a). Signs of deficiency may occur rather quickly if
dietary phosphorus is insufficient. Deficiency is most com-
mon in cattle grazing forages on soils low in phosphorus
or in animals consuming excessively mature forages or crop
residues with low phosphorus content (less than 0.25 per-
cent, dry basis). Nonspecific chronic signs of deficiency
include unthriftiness, inappetence, poor growth and lacta-
tional performance, and unsatisfactory fertility; but signs
are often complicated by coincidental deficiencies of other
nutrients such as protein or energy. Animals maybe chroni-
cally hypophosphatemic (low phosphorus in blood
plasma 2 to 3.5 mg/dl), but the concentration of phospho-
rus in milk remains within the normal range. In severe
deficiency cases, bone mineral mass is lost, and bones
OCR for page 151
Minerals 151
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Representative terms from entire chapter:
dairy cattle
